Howard J. Falcon-Lang

Pennsylvanian tropical rain forests responded to glacial-interglacial rhythms

Pennsylvanian tropical rain forests flourished during an icehouse climate mode. Although it is well established that Milankovitch-band glacial-interglacial rhythms caused marked synchronous changes in Pennsylvanian tropical climate and sea level, little is known of vegetation response to orbital forcing. This knowledge gap has now been addressed through sequence- stratigraphic analysis of megafloral and palynofloral assemblages within the Westphalian D–Cantabrian Sydney Mines Formation of eastern Canada. This succession was deposited in a low- accommodation setting where sequences can be attributed confidently to glacio-eustasy. Results show that long-lived, low-diversity peat mires dominated by lycopsids were initiated during deglaciation events, but were mostly drowned by rising sea level at maximum interglacial conditions. Only upland coniferopsid forests survived flooding without significant disturbance. Mid- to late interglacial phases witnessed delta-plain progradation and establishment of high-diversity, mineral-substrate rain forests containing lycopsids, sphenopsids, pteridosperms, cordaites, and tree ferns. Renewed glaciation resulted in sea-level fall, paleovalley incision, and the onset of climatic aridity. Glacial vegetation was dominated by cordaites, pteridosperms, and tree ferns; hydrophilic lycopsids and sphenopsids survived in paleovalley refugia. Findings clearly demonstrate the dynamic nature of Pennsylvanian tropical ecosystems and are timely given current debates about the impact of Quaternary glacial-interglacial rhythms on the biogeography of tropical rain forest.

Incised channel fills containing conifers indicate that seasonally dry vegetation dominated Pennsylvanian tropical lowlands

The idea that the Pennsylvanian tropical lowlands were temporally dominated by rainforest (i.e., the Coal Forest) is deeply ingrained in the literature. Here we challenge two centuries of research by suggesting that this concept is based on a taphonomic artifact, and that seasonally dry vegetation dominated instead. This controversial finding arises from the discovery of a new middle Pennsylvanian (Moscovian) fossil plant assemblage in southeast Illinois, United States. The assemblage, which contains xerophytic walchian conifers, occurs in channels incised into a calcic Vertisol below the Baker Coal. These plants grew on seasonally dry tropical lowlands inferred to have developed during a glacial phase. This xerophytic flora differs markedly from that of the typical clubmoss-dominated Coal Forest developed during deglaciation events. Although preserved only very rarely, we argue that such xerophytic floras were temporally as dominant, and perhaps more dominant, than the iconic Coal Forests, which are overrepresented in the fossil record due to taphonomic megabias. These findings require the iconography of Pennsylvanian tropical lowlands to be redrawn.

Rainforest collapse triggered Carboniferous tetrapod diversification in Euramerica

Abrupt collapse of the tropical rainforest biome (Coal Forests) drove rapid diversification of Carboniferous tetrapods (amphibians and reptiles) in Euramerica. This finding is based on analysis of global and alpha diversity databases in a precise geologic context. From Visean to Moscovian time, both diversity measures steadily increased, but following rainforest collapse in earliest Kasimovian time (ca. 305 Ma), tetrapod extinction rate peaked, alpha diversity imploded, and endemism developed for the first time. Analysis of ecological diversity shows that rainforest collapse was also accompanied by acquisition of new feeding strategies (predators, herbivores), consistent with tetrapod adaptation to the effects of habitat fragmentation and resource restriction. Effects on amphibians were particularly devastating, while amniotes (‘reptiles’) fared better, being ecologically adapted to the drier conditions that followed. Our results demonstrate, for the first time, that Coal Forest fragmentation influenced profoundly the ecology and evolution of terrestrial fauna in tropical Euramerica, and illustrate the tight coupling that existed between vegetation, climate, and trophic webs.

The Pennsylvanian tropical biome reconstructed from the Joggins Formation of Nova Scotia, Canada

The Pennsylvanian (Langsettian) Joggins Formation contains a diverse fossil assemblage, first made famous by Lyell and Dawson in the mid-19th century. Collector curves based on c. 150 years of observation suggest that the Joggins fossil record is relatively complete. A key feature of the site is that fossils occur in (par)autochthonous assemblages within a narrow time interval (<1 Ma). Analysis of co-occurring taxa within a precise facies context permits ecosystem reconstruction, and three main communities are recognized in this study. Brackish seas, the distal extension of European marine bands, were populated by Foraminifera, molluscs, annelids, arthropods, fishes, and aquatic tetrapods. Poorly drained coastal plains were covered by rainforests of lycopsids, calamiteans, ferns, pteridosperms, and cordaitaleans, inhabited by a terrestrial fauna of molluscs, annelids, arthropods, and tetrapods, including the earliest known reptiles. Well-drained alluvial plains were covered by fire-prone cordaitalean scrub containing a low-diversity fauna of molluscs, arthropods, and tetrapods, locally preserved in waterholes. These three environments repeatedly interchanged with one another in response to base-level fluctuations forced by tectonism and glacioeustasy. Located further inland than other well-studied Pennsylvanian tropical sites, the Joggins Formation is significant because it contains a record of intra-continental terrestrial ecosystems.

Pennsylvanian uplands were forested by giant cordaitalean trees

The precise timing of when upland terrains first became forested is highly controversial. Pennsylvanian palynoflora and megaflora transported into marine highstand deposits imply that emergent topographic highs may have supported cordaitalean forests. The discovery of a new Pennsylvanian (Bolsovian) plant assemblage in southwest Newfoundland confirms this hypothesis and allows the architecture of these upland trees to be reconstructed in detail. The assemblage includes several hundred calcareously permineralized stumps, trunks, and branches, and represents the remains of shallowly rooted cordaitalean trees that were ≤48.5 m high when mature. The fossils occur in alluvial conglomerates that constitute a 10-km-diameter outlier on the margins of the paleoequatorial Variscan foreland. The paleogeographic setting together with plant taphonomic inferences strongly indicate that these giant trees were transported from nearby upland alluvial plains and deposited in an elevated intermontane basin. This interpretation is supported by analysis of rootstock morphology, which implies tree growth in thin soils consistent with an alluvial gravel substrate. This improved understanding of Pennsylvanian upland forests has important implications for geochemical modeling of the global carbon cycle.

What happened to the coal forests during Pennsylvanian glacial phases

Abstract Sequence stratigraphic analysis of Pennsylvanian coal-bearing strata suggests that glacial-interglacial fluctuations at high latitudes drove cyclic changes in tropical biomes. A literature review of plant assemblages in this paleoclimatic context suggests that coal forests dominated during humid interglacial phases, but were replaced by seasonally dry vegetation during glacial phases. After each glacial event, coal forests reassembled with largely the same species composition. This remarkable stasis implies that coal-forest refugia existed across the equatorial landscape during glacial phases, expanding to repopulate lowlands during and following deglaciation. One possibility is that refugia comprised small pockets of wetland forest strung out along valleys at some sites, but data are currently insufficient to test this hypothesis. The model presented here, if accepted, dramatically alters our understanding of the coal forests and helps explain aspects of their dynamics.

Biodiversity and terrestrial ecology of a mid-Cretaceous, high-latitude floodplain, Alexander Island, Antarctica

The biodiversity and terrestrial ecology of the Late Albian Triton Point Formation (Fossil Bluff Group), Alexander Island, Antarctica is analysed to improve our understanding of polar biomes during the mid-Cretaceous thermal optimum. This formation was deposited on a high-latitude (75°S) floodplain and consists of two facies associations, a lower braided alluvial plain unit and an upper coastal meander-belt unit. Analysis of fossil plants in well exposed palaeosols reveals the existence of spatially complex plant communities. Braidplains supported patchy, low-density (91 trees/ha) stands of podocarp and taxodioid conifers on floodbasin substrates, and conifer–cycadophyte–fern–angiosperm thickets in riparian settings. Coastal meander-belts supported medium density (568 trees/ha) podocarp–araucarian conifer forests on mature floodbasin soils, and fern–angiosperm–ginkgo thickets in riparian settings. Growth-ring analysis indicates plants experienced stressful growing conditions on the braidplain characterized by high-frequency flood events, but more favourable growing conditions on the coastal plain. Additional vegetation disturbances were caused by arthropod–fungal attack, frost and wildfire. In terms of structure, composition, ecology and productivity these predominantly evergreen, broad-leafed conifer forests bear similarities to the extant temperate rainforests of New Zealand.

Cretaceous (Late Albian) coniferales of Alexander Island, Antarctica. 1: Wood taxonomy: a quantitative approach

Silicified conifer woods are very common in the mid-Cretaceous (Late Albian, 100Ma) Triton Point Member of the Neptune Glacier Formation (Fossil Bluff Group), SE Alexander Island, Antarctica. These occur as up to 7m high in situ tree trunks and stumps rooted in carbonaceous palaeosols and as allochthonous logs and wood fragments in fluvial channel and sheet sandstone facies. Sixty-eight wood samples were examined in this study and were classified in terms of five form taxa using a quantitative approach. Araucarioxylon (1.5% of specimens) is characterised by dominantly multiseriate, alternately arranged bordered pitting on radial tracheid walls and by 1-4 araucarioid cross-field pitting. Araucariopitys (11.8% of specimens) is characterised by dominantly uniseriate tracheid pitting with subordinate biseriate, alternate tracheid pitting and by 1-4 araucarioid cross-field pitting. Podocarpoxylon sp. 1 (63.1% of specimens) is characterised by contiguous, uniseriate tracheid pitting and 1-2 podocarpoid cross-field pits. Podocarpoxylon sp. 2 (22.1% of specimens) is similar to P. sp. 1, differing only in that ray height is lower, tracheid pits are dominantly spaced more than one pit diameter apart and abundant axial parenchyma is present. These first four taxa all possess growth rings with subtle boundaries. Taxodioxylon (1.5% of specimens) is characterised by 1-2 seriate, oppositely arranged, bordered tracheid pitting, 1-2 taxodioid cross-field pitting and very marked ring boundaries. These woods were derived from large trees with basal stump diameters of up to 0.5m and probable heights of up to 29m. Data from leaf traces suggest that Araucariopitys and Podocarpoxylon sp. 1 and sp. 2 (97% of specimens) were evergreen with leaf retention times of >5years. These predominantly evergreen conifer forests grew in a mild, high latitude (75 degrees S) environment during the mid-Cretaceous greenhouse climate phase.

Late Carboniferous Tropical Dryland Vegetation in an Alluvial-plain Setting, Joggins, Nova Scotia, Canada

Abstract The composition and ecology of Late Carboniferous tropical, dryland plant communities are described for the first time. The study focuses on a 700-m-thick succession through the Langsettian Joggins Formation. Red bed units (5–110 m thick) bearing isolated pedogenic carbonate nodules occur at fourteen intervals and are interpreted as originating in a seasonally dry, well-drained, alluvial-plain setting characterized, in places, by an anastomosing fluvial geometry. A quantitative quadrat analysis of red bed floral assemblages preserved as compressions, impressions, calcareous permineralizations, and charcoal was undertaken. Cordaites, represented by woody trunks, branches, pith casts, leaves, and seeds, comprise 74% of the red bed floral thanatomass, together with minor pteridosperms and sphenopsids, and rare sigillarian lycopsids. Taphonomic data demonstrate that while fire-prone cordaite-pteridosperm vegetation dominated nearly all seasonally dry floodplain niches, hydrophilic lycopsids and sphenopsids were restricted to riparian settings where water availability was greatest. Calculation of standard diversity indices indicates that growing conditions were generally stressful, consistent with a seasonal environment. The composition of these dryland communities differs markedly from lycopsid-dominated wetland communities known from gray, coal-bearing successions at other intervals in the Joggins Formation. Sequence stratigraphic analysis of the Joggins Formation suggests that dryland communities represent the vegetation of seasonal continental-interior environments in contrast to wetland communities that grew in humid coastal settings. Repeated alternations between continental and coastal settings were caused by tectonically and/or eustatically driven base-level fluctuations that resulted in marine transgressive-regressive rhythms minimally hundreds of kilometers long.

Ecological Gradients within a Pennsylvanian Mire Forest

Pennsylvanian coals represent remains of the earliest peat-forming rain forests, but there is no current consensus on forest ecology. Localized studies of fossil forests suggest intermixture of taxa (heterogeneity), while, in contrast, coal ball and palynological analyses imply the existence of pronounced ecological gradients. Here, we report the discovery of a spectacular fossil forest preserved over ∼1000 ha on top of the Pennsylvanian (Desmoinesian) Herrin (No. 6) Coal of Illinois, United States. The forest was abruptly drowned when fault movement dropped a segment of coastal mire below sea level. In the largest study of its kind to date, forest composition is statistically analyzed within a well-constrained paleogeographic context. Findings resolve apparent conflicts in models of Pennsylvanian mire ecology by confirming the existence of forest heterogeneity at the local scale, while additionally demonstrating the emergence of ecological gradients at landscape scale.